EDINB. J. BOT.

48(3): 307-332 (1991)

A COMPARATIVE STUDY OF FIVE CERRADO AREAS IN

SOUTHERN MATO GROSSO, BRAZIL
A. T. OLIVEIRA-FILHO* & F. R. MARTINS**
Quantitative descriptions of the woody flora and soil analyses are given for five areas of
cerrado (savanna woodland) near Cuiaba, Mato Grosso state, Brazil. Comparisons are made
of the floristic composition and community structure of these areas using polar ordination and
hierarchical classification techniques.
Descricoes quantitativas da flora lenhosa e analises de solos sao apresentadas para cinco areas
de cerrado nas proximidades de Cuiaba, Mato Grosso. Sao realizadas comparacoes entre estas
areas em composicao floristica e estrutura comunitaria utilizando tecnicas de ordenacao polar
e classificacao hierarquica.

Cerrado is a savanna-like vegetation which covers about 2 million km , representing

about one-fifth of Brazil's total area. In the state of Mato Grosso, it occupies about 28.5%
(251085km2) of the total area and abuts on the Amazonian forest (Hylaea) to the north
and the Pantanal Matogrossense (the low-lying floodplain of the Rio Paraguai) to the
south (Fig. 1). It is currently being destroyed very rapidly: in 1980, 12% or 30069km
of the Mato Grosso cerrado was already completely cleared for cattle-pasture and crops,
particularly soya, and since then this area has probably more than doubled (Cameiro,
1982). The floristic knowledge of this vanishing vegetation is very incomplete. Few
surveys of Mato Grosso cerrados are available in the literature. A number of
taxonomists who participated in the first scientific expeditions to Mato Grosso (from
1790 to 1915) left some general descriptions and checklists for the regional flora (see
Hoehne, 1923 for a review). Pioneer detailed studies of particular areas of cerrado were
carried out by Ratter (1971), Ratter et al. (1973, 1977) and Furley et al. (1988) in
northeastern Mato Grosso. Oliveira-Filho & Martins (1986) and Oliveira-Filho et al.
(1989) provided floristic and phytosociological surveys of cerrados of the region of
Cuiaba, the capital of the state. For the seasonally flooded cerrados of the Pantanal
Matogrossense, Prance & Schaller (1982) gave a detailed floristic survey while Pott et
al. (1986) contributed a detailed checklist and Ratter et al. (1988) a quantitative-floristic
study of an important area.
The present communication is devoted to a quantitative study of the woody flora of
five cerrado areas, on different soils, in the region of Cuiaba and the Chapada dos
Guimaraes, in central-southern Mato Grosso. Comparisons are made to evaluate dif-

Departamento de Ciecias Florestais, Escola Superior de Agricultura de Lavras, 37200

Lavras, MG, Brazil
** Departamento de Botanica, Instituto de Biologia, Universidade Estadual de Campinas,
13081 Campinas, SP, Brazil.

307

EDINB. J. BOT. 48(3)

308

".^;i;.<.;..-.' GUIMARAES-.;..,
> '

'"' A

'

'

"

:>;..".. .-foChapada dos .' . ." /

.' /''i,,Guimaraes. '. r

Fazenda
Sao Pedro
Bandeira
Km 14

BAIXADA
CUIABANA
Sto. Antonio
do
Leverger

56 30'

56 00'

5530'

FIG. 1. Map of the region of Cuiaba and Chapada dos Guimaraes showing the three study areas. The dotted area
is above 400m in altitude; dense supple = town; BR = federal road; MT = state road.

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

309

ferences in floristic composition and community structure of these areas related to

variations in soil properties.
DESCRIPTION OF THE STUDY AREAS
The region of Cuiaba and the Chapada dos Guimaraes comprises a wide low plain, the
Baixada Cuiabana, and a higher plateau, the Chapada dos Guimaraes (Fig. 1). The
Baixada Cuiabana is a flattish surface with altitudes ranging from c.lOOm, in the
southern border with the Pantanal Matogrossense, to c.350m, at the base of the Chapada
dos Guimaraes. This low plateau is the western tip of a large area of tablelands: the
Planalto Central. The Chapada dos Guimaraes is bounded by vertical escarpments of
5O-3OOm and the relief at the top is relatively flat with altitudes ranging from 500 to
800m. This high plateau is composed of two layers of Devonian sandstone which overlie
the Pre-Cambrian phyllite of the Baixada Cuiabana.
The climate is tropical and continental with high temperatures all year round and a
rainy summer and dry winter, being classified as Koppen's 'Savanna Subtype' (Aw) of
the tropical rain climate (A) (Eidt, 1968). Detailed climatic data are available only for
Cuiaba where the mean annual air temperature is 25.6C, with monthly means ranging
from 22.8C (July) to 27.2C (October), and the mean annual rainfall of about 1420mm
is concentrated during October-May (95% of the total rainfall) (Bombled, s.d.). The
temperatures of the Chapada dos Guimaraes, however, are 3 ^ C lower than Cuiaba and
the annual rainfall can reach 2000mm (EDIBAP, 1979).
The main vegetation type of the region is cerrado with its characteristic wide physiognomic variations. Forests are mostly restricted to galleries along the watercourses and
to some patches at the top of the high plateau. Grasslands can be found in rocky and/or
in poorly-drained sites. The vegetation is relatively well-preserved considering that the
colonization of the region by Europeans goes back to the beginning of the 18th century:
the very infertile soils have restricted the spread of cultivated areas throughout the region
and human damage is caused principally by repeated fires and more localized gold
mining and urban growth. In 1989 a part of the Chapada dos Guimaraes was created a
National Park.
The five areas of cerrado studied lie in three sites along the road (the MT 305) which
connects Cuiaba to the Chapada dos Guimaraes: Fazenda Sao Pedro-Bandeira (km 14)*,
the Rio Claro region (km 42) and the Salgadeira region (km 45) (see Fig. 1).
Site A: Fazenda Sao Pedro-Bandeira (1530'S, 5602'W, c.l90m alt.) This is a ranch
where the cerrado is used as natural pasture for cattle at low density. Two areas were
studied here:
Area 1 - Murundu cerrado: This type of cerrado occurs on round earthmounds (0.220.0m diameter, 0.1-2.0m high and usually bearing termitaria) scattered in a regular

* The kilometre numbers refer to the distance from Cuiaba. They are marked by roadsigns.

aip pus ( H D ) OSSOJQ OJEJAJ sp IBiapaj apEpisjSAiun jo BUEqjsq aqj ui paSpoj are
suauipads isipnoA 'siured puB (sasxi pus sqnjqs 'sqtuqsqns) sapads XpooA\ 01 USAIS
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uonBj93A aqj jo XuiouSoisXqd aqj, opBJjaojo spuBjsi anji SE snputunui 9ip SUIAESI ESJB
PUEJSSBJS 9joqA\ aqj SJ3A00 daap uiooi "0 JSXBJ B uaqM 'UOSBSS XUIBJ aqj guunp uiqiiM
J91BAV aqj urep jEqj sisajo X>JOOJ Xq papunoxms (Bq90"5) uoissajdap IBIAIIIIB JEU E UI jnooo
BJiapuBa-ojpaj OBS npuazEj jo snpuiunui aqx (punoujqiiEa = nputunui 'spuBjssEJS =
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'ioa

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

311

Universidade Estadual de Campinas (UEC) with duplicates at the Escola Superior de

Agricultura de Lavras (ESAL) and the Royal Botanic Garden, Edinburgh (E). A list of
all species recorded is given in the Appendix.
In each of the five study areas, a number of 0.5 litre samples of soil were collected
from the depth of 0-20cm and mixed up to form a compound sample. Chemical and
granulometric analyses were made at the Soil Laboratory of the Universidade Federal
de Mato Grosso.
Woody plants were sampled using the point-centred quarter method (PCQ), as
described by Mueller-Dombois & Ellenberg (1974), in three areas of cerrado: the pebble
cerrado (Area 2) with 140 points distributed in a 4 x 35 systematic grid, the phyllite
cerrado (Area 3) with 60 points in a 5 x 12 grid, and the sandstone cerrado (Area 5) with
50 points in two 5 x 5 grids. Inter-point distances were of 10m. The interfluvial cerrado
(Area 4) was sampled by 100 contiguous plots of 10 x 10m, arranged in a transect of 50
paired plots. The murundu cerrado (Area 1) was sampled on 80 earthmounds (0.5-22m
diameter), comprising in total 0.3ha of murundus. In all surveys, species, height, and
circumference at the stem base were recorded for all shrubs, trees and palms with a stem
circumference of at least 9cm. The number of individuals, relative density, total basal
area, relative dominance (based on basal area), relative frequency, importance value
index (IVI = rel.dens. + rel.dom. + rel.freq.), and cover value index (CVI = rel.dens. +
rel.dom.) were calculated for each species. The CVI was also calculated for each plant
family. The Shannon-Wiener diversity index and the Pielou evenness index (Brower &
Zar, 1984) were obtained for each sampled community.
The five cerrado communities were compared using three dissimilarity measures: the
complemented coefficient of community (CC), the percentage difference (PD) and the
Euclidian distance (ED), following Gauch (1982):

cc,, = ioo-

200sc

200.

PDp=100-

where si and S2 = number of species in communities 1 and 2; sc = number

of species in common in the two communities; An, Ai2 = abundance of
species i in communities 1 and 2; min [An, Ai2] = lowest value of
abundance of species i in communities 1 and 2.
The abundance values used for the species in the calculations of PDs and EDs were their
CVIs. ED is a measure determined mostly by larger abundance values while CC is
determined only by the floristic composition. PD is intermediate, with linear weighting
of species (Gauch, 1982). The dissimilarity matrices of CCs, PDs and EDs were used to

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JO 9jnjB9J SupfUlS 1S0UI aqi 'J9A9MOH '(/, 3IQEX) IV J 90IHA J9M0J aqi Xq pUB 'S1U3JUO0
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puB /?u ;y(Ct/50yi 'vuopouvjjVAUOo DJU3AJVS 'vjojfituvd v9jvnQ - XJIUIBJ siqi jo sspads
moj :( Sij) IAD uj ^I?urej juBjioduii JSOUI aqi SJB 9B30BTsXqooA aqi 'opBiiao nputunm
SuunoqqSiau aqi oj ISBJIUOO U\ (9 ajqBx) jaiauiBip puB jqBpq SSBJOAB JO ssnjBA
SB j p M SB aouBUTtuop pus Xjisuap ain]osqB jo sanjBA J3/v\0] ssq opBxiao ajqqad

U
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^ pus iv J3qSiq 'Hd J9A\OJ Xq pazuaiOBJBqo SIB snpuainui aqi jo SJIOS sqx '(Z g H
opBJjao npuainui aqi ui juBiiodunun XiqBjou 3JB 'sopBxiao jaqjo aqi ui I A D
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aan JJBJ OI MOI B SB Xpuanbaij ssaj puB ( U I ^ - I ) qruqs pajiuaqninui B SB XIUIBUI SJTIODO
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vaunv vmqdqvi puB sisudqv&no vjjpuy 's)\npa vijj9qj]V 'vn>p xLiaidiQ saioads
sapads i9A\aj in siBnpiAipui jo uoqBjjuaouoo jo aaritep aqi samsBaui qo]qA\ (9
'f) ssauuaAa jo aniBA jssMOi aqi SuiABq opexiao npumntu aqi JOJ UOSBSJ aqi jo qonui
suiBidxa OSIB aouBuiuiopaid SJI 1 A I I^ioi jo %pp JOJ aiqisuodsai Supq 'Xouanbaij pire
aouBuiuiop 'Ajisuap jo sanreA jsaqSiq aqi JBJ Xq qiiM sapads aqi si vuvouautv vjpjDunj
siBnpiAipui pajduiBS 09S ui sapads 59 qiiA\ 'opBjjao aiqqad SuijnoqqSpu aqi ireqj
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puB 'aouBuiuiop pus Xjisuap ajnjosqB IBJOJ JO sanreA jsaqSiq ai^ siuasajd 11 :srenpiAipui
jreuis snojauinu jo aouBuiuiop-oo aqi Xq pazuajoBJBqo si opBJjao npurunui aqx
suiBjgojpuap XiuBiiuiissip uo puB SUIBJSEIP uopBuipjo
jB]od uo sapiunuiuioo opsjjao aqi jo SISXJBUB aAUBJBduioo aqj siuasajd -gij /, ajqBx ui
uMoqs aJB sasXreuB JIOS jo sjjnsaj aqx 'Z '%}d u? parexisnui ajB saijiurej iirejjoduii JSOUI
aqi jo (SJAD) saoipui an^BA J3AOO aqx 9 ^iq^x ui uaAiS ajB Xiiuntutuoo qoEa JOJ BIBP
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suiBjSojpuap XiuBiiuiissip puB (3861 'M 3nB 9) SUIBJSBIP uorreuipjo jBjod qioq ajBdajd

10H 'f

A COMPARATIVE STUDY OF CERR ADO AREAS IN BRAZIL

313

TABLE 1. Murundu cerrado: species sampled on 80 earthmounds (0.5-20.0m in

diameter; 0.321 ha in total area). n = number of individuals; ba = basal area (m2);
Rel.dens. = relative density; Rel.dom. = relative dominance; Rel.freq. = relative
frequency; IVI = importance value index. Species in descending order of IVI.
Total
ba

n
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.

Curatella americana
Dipteryx alata
Alibertia edulis
Andira cuyabensis
Tabebuia aurca
Hymenaea stigonocarpa
Eugenia aurata
Erythroxylum suberosum
Pseudobombax longiflorum
Simarouba vcrsicolor
Vatairea macrocarpa
Matayba guianensis
Tabebuia caraiba
Sclerolobium aureum
Byrsonima pachyphylla
Jacaranda cuspidifolia
Astronium fraxinifolium
Abuta selloana
Cecropia pachystachya
Brosimum gaudichaudii
Copaifera martii
Pouteria ramiflora
Diospyros coccolobifolia
Cybistax antisyphilitica
Bowdichia major
Casearia sylvestris
Heteropterys byrsonimifolia
Qualea parviflora
Campomanesia eugenioides
Erythroxylum citrifolium
Simaba trichilioides
Zanthoxylum cinereum
Bauhinia bongardi
Myrcia formosiana
Himatanthus obovatus
Magonia pubescens
Rudgea viburnioides
Kielmeyera coriacea
Vernonia brasiliana
Ouratea hexaspcrma
Myrcia uberavensis
Cardiopetalum calophyllum
Connarus suberosus
Annona pygmaea
Tocoyena formosa

39C
At
8:
4<:
y.
l:
3:
IS

Totals

871

r.
I.5
>

rl.l!
i
i>
>
1()
ii
7
ii
5
4
4
5
(5
4
;5
5
1
I
>

Re
dens.

Rel.
dom.

Rel.
freq.

IVI

4.194
2.053
0.471
0.503
0.479
0.777
0.135
0.112
0.078
0.225
0.273
0.022
0.093
0.047
0.054
0.145
0.053
0.066
0.142
0.025
0.047
0.040
0.036
0.021
0.028
0.022
0.023
0.014
0.013
0.002
0.002
0.031
0.003
0.008
0.004
0.004
0.003
0.002
0.002
0.002
0.001
0.001
0.001
0.001
0.001

45.46
5.28
9.41
5.05
4.02
1.72
3.79
3.33
2.64
1.72
0.92
1.95
1.49
0.92
1.03
0.57
1.15
0.92
0.80
0.92
0.57
0.46
0.46
0.57
0.69
0.46
0.34
0.34
0.80
0.23
0.23
0.11
0.23
0.1 1
0.11
0.1 1
0.11
0.1 1
0.1 1
0.1 1
0.1 1
0.1 1
0.1 1
0.1 1
0.1 1

40.89
20.02
4.59
4.91
4.67
7.57
1.32
1.10
0.76
2.20
2.66
0.21
0.91
0.45
0.53
1.41
0.52
0.64
1.39
0.24
0.46
0.39
0.35
0.21
0.27
0.22
0.23
0.13
0.13
0.02
0.02
0.30
0.03
0.08
0.04
0.03
0.03
0.02
0.02
0.02
0.01
0.01
0.01
0.01
0.01

16.92
8.33
8.84
8.08
6.06
2.27
5.30
5.81
4.29
3.54
2.02
3.28
2.78
2.02
.77
.26
.52
.52
).76
.77
.26
.01
.01
.01
0.51
0.76
0.76
0.76
0.25
0.51
0.51
0.25
0.25
0.25
0.25
0.25
0.25
0.25
0.25
0.25
0.25
0.25
0.25
0.25
3.25

103.27
33.63
22.84
18.04
14.75
11.57
10.41
10.23
7.69
7.45
5.60
5.44
5.18
3.39
3.33
3.25
3.18
3.08
2.95
2.93
2.29
.86
.82
.79
.46
.44
.33
.23
.19
0.75
0.75
0.67
0.51
0.45
0.40
0.40
0.40
0.39
0.39
0.38
0.38
0.38
0.38
0.38
3.38

10.258

100

100

100

300

EDINB. J. BOT. 48(3)

314

TABLE 2. Pebble cerrado: species sampled by point-centred quarter method in a

systematic grid of 140 points. See Table 1 for abbreviations.
n
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.

Qualea parviflora
Curatella americana
Salvertia convallariodora
Myrcia uberavensis
Vochysia rufa
Qualea grandiflora
Dipteryx alata
Pouteria ramiflora
Salacia crassifolia
Da villa elliptica
Bowdichia major
Pseudobombax longiflorum
Connarus suberosus
Andira cuyabensis
Tabebuia caraiba
Sclerolobium aureum
Byrsonima coccolobifolia
Byrsonima pachyphylla
Simarouba versicolor
Vatairea macrocarpa
Erythroxylum suberosum
Brosimum gaudichaudii
Diospyros coccolobifolia
Astronium fraxinifolium
Hymenaea stigonocarpa
Dimorphandra mollis
Tabebuia ochracea
Tabebuia aurea
Ouratea spectabilis
Annona coriacea
Himatanthus obovata
Lafoensia pacari
Agonandra brasiliensis
Magonia pubescens
Ouratea hexasperma
Ferdinandus elliptica
Strychnos pseudoquina
Mataybaguianensis
Dilodendron bipinnatum
Eriotheca gracilipes
Pseudobombax tomentosum
Myrcia pallens
Kielmeyera coriacea
Diptychandra aurantiaca
Kielmeyera rubriflora
Acrocomia aculeata
Dalbergia miscolobium
Jacaranda cuspidifolia
Platypodium elegans
Aspidosperma macrocarpon
Alibertia edulis
Hancornia speciosa
Eugenia aurata
Stryphnodendroii obovatum
Hirtella gracilipes
Rudgea viburnioides

81
53
48
42
27
26
13
19
17
17
11
9
15
9
10
12
8
8
7
4
8
9
9
6
4
6
6
6
5
5
4
4
4
3
3
3
2
3
2
2
2
2
2
2
2

Total
ba

Rel.
dens.

Rel.
dom.

Rel.
freq.

1.212
1.086
0.663
0.303
0.645
0.384
0.599
3.325
0.209
0.116
0.194
0.219
0.054
0.171
0.134
0.027
0.124
0.098
0.114
0.197
0.063
0.033
0.024
0.087
0.145
0.073
0.058
0.051
0.042
0.013
0.039
0.026
0.023
0.053
0.034
0.029
0.053
0.015
0.026
0.020
0.013
0.009
0.008
0.005
0.005
0.034
0.030
0.022
0.022
0.018
0.011
0.010
0.008
0.006
0.006
0.005

14.46
9.46
8.57
7.50
4.82
4.64
2.32
3.39
3.04
3.04
.96
.61
;'..68
.61
.79
>.14
.43
.43
.25
().71
.43
.61
.61
.07
().71
.07
.07
.07
0.89
0.89
0.71
0.71
0.71
0.54
0.54
0.54
0.36
0.54
0.36
0.36
0.36
0.36
0.36
0.36
0.36
0.18
0.18
0.18
0.18
0.18
0.18
0.18
0.18
0.18
0.18
C).18

15.11
13.55
8.27
3.78
8.04
4.79
7.47
4.06
2.61
1.45
2.42
2.73
0.68
2.13
1.70
0.34
1.54
1.22
1.42
2.46
0.79
0.41
0.30
1.09
1.80
0.91
0.73
0.63
0.53
0.17
0.49
0.33
0.29
0.66
0.42
0.36
0.66
0.19
0.32
0.25
0.16
0.11
0.09
0.06
0.06
0.42
0.37
0.28
0.27
0.22
0.14
0.12
0.10
0.08
0.07
0.06

13.15
9.36
7.57
6.97
4.18
4.78
1.99
3.59
3.19
2.99
2.19
1.79
2.39
.79
.99
;1.39
.59
.59
.39
().8O
.59
.59
.59
.20
().8O
.20
.20
.20
.00
.00
0.80
0.80
0.80
0.60
0.60
0.60
0.40
0.60
0.40
0.40
0.40
0.40
0.40
0.40
0.40
0.20
0.20
0.20
0.20
0.20
0.20
0.20
0.20
0.20
0.20
0.20

IVI
42.73
32.37
24.42
18.25
17.04
14.22
11.79
11.04
8.83
7.47
6.57
6.13
5.75
5.54
5.48
4.87
4.56
4.24
4.07
3.97
3.81
3.61
3.50
3.36
3.31
3.18
2.99
2.90
2.42
2.05
2.00
.84
.80
.80
.56
.50
.42
.32
.07
.01
0.91
0.87
0.85
0.85
0.82
0.80
0.75
0.66
0.65
0.60
0.51
0.50
0.48
0.46
0.45
0.45

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

315

TABLE2(Cont.)
Total
ba
57.
58.
59.
60.
61.
62.
63.
64.
65.

0.005
0.004
0.003
0.003
0.003
0.002
0.002
0.002
0.001

Coccoloba mollis
Eugenia uniflora
Neca theif'era
Coussarca hydrangcaefolia
Cardiopctalum calophyllum
Cybistax antisyphililica
Acosmium dasycarpum
Casearia sylvestris
Tocoycna formosa
Totals

560

8.018

Rel.
dens.
0.18
0.18
0.18
0.18
0.18
0.18
0.18
0.18
0.18
100

Rel.
dom.
0.06
0.05
0.04
0.04
0.03
0.03
0.02
0.02
0.01
100

Rel.
freq.
0.20
0.20
0.20
0.20
0.20
0.20
0.20
0.20
0.20
100

IVI
0.43
0.43
0.42
0.41
0.41
0.40
0.40
0.40
0.39
300

TABLE 3. Phyllite cerrado: species sampled by point-centred quarter method in a

systematic grid of 60 points. See Table 1 for abbreviations.

1.
2
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.

Magonia pubesccns
Callisthenc fasciculata
Pscudobombax longiflorum
Dipteryx alala
Astronium fraxinifolium
Vatairea macrocarpa
Jacaranda cuspidifolia
Anadenanthera peregrina
var. falcata
Curatella americana
Terminalia subsericea
Lafocnsia pacari
Austroplenckia populnca
Byrsonima pachyphylla
Qualea parviflora
Bowdichia major
Byrsonima coccolobifolia
Kielmcyera rubriflora
Sclerolobium aurcum
Dimorphandra mollis
Acrocomia aculeata
Campomanesia eugenioides
Eriotheca gracilipes
Aspidosperma nobile
Hymenaea sligonocarpa
Qualea multiflora
Machaerium acutifolium
Kielmeyera coriacea
Tabebuiaochracea
Guettarda viburnioides
Myrcia uberavensis
Himatanthus obovatus
Connarus suberosus
Luchea speciosa
Acosmium dasycarpum
Hancornia speciosa
Plathvmenia rcticulata

Total
ba

Rel.
dens.

Rel.
dom.

Rel.
freq.

IVI

32
22
20
11
10
11
7

0.355
0.330
0.124
0.254
0.215
0.111
0.185

13.33
9.17
8.33
4.58
4.17
4.58
2.92

14.18
13.20
4.94
10.16
8.60
4.42
7.38

11.98
7.83
8.76
4.15
4.61
4.61
3.23

39.50
30.20
22.03
18.89
17.37
13.62
13.53

8
9
11
8
5
7
6
5
5
5
4
4

0.135
0.090
0.031
0.033
0.078
0.026
0.029
0.036
0.024
0.020
0.038
0.022
0.032
0.031
0.021
0.021
0.020
0.020
0.041
0.008
0.010
0.018
0.010
0.009
0.006
0.023
0.015
0.015
0.010

3.33
3.75
4.58
3.33
2.08
2.92
2.50
2.08
2.08
2.08
1.67
1.67
1.25
1.25
1.25
1.25
1.25
1.25
0.83
1.25
1.25
0.83
0.83
0.83
0.83
0.42
0.42
0.42
0.42

5.40
3.62
1.25
1.33
3.11
1.03
1.16
1.42
0.98
0.81
1.54
0.89
1.26
1.25
0.84
0.82
0.81
0.81
1.64
0.30
0.41
0.72
0.40
0.36
0.22
0.93
0.59
0.59
0.39

3.23
3.69
3.69
3.23
2.30
3.23
2.30
2.30
2.30
2.30
.84
.84
.38
.38
.38
.38
.38
1.38
0.92
1.38
0.92
0.92
0.92
0.92
0.92
0.46
0.46
0.46
).46

11.95
11.05
9.52
7.89
7.49
7.17
5.96
5.81
5.37
5.20
5.05
4.40
3.90
3.88
3.48
3.46
3.44
3.44
3.40
2.94
2.59
2.48
2.16
2.11
1.97
1.81
1.47
1.47
1.27

3
3
3
3
3
3
2
3
3
2
z

2
2
1
1
1
1

EDINB. J. BOT. 48(3)

316

TABLE3(Cont.)
Total
ba

Rel.
dens.

Rel.
dom.

1
1
1
1
1
1
1
1
1
1
1
1

0.008
0.007
0.007
0.005
0.005
0.005
0.005
0.005
0.003
0.003
0.002
0.001

0.42
0.42
0.42
0.42
0.42
0.42
0.42
0.42
0.42
0.42
0.42
0.42

0.31
0.29
0.27
0.22
0.20
0.20
0.20
0.18
0.11
0.11
0.09
0.05

240

2.501

n
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.

Peltogyne confertiflora
Pouteria ramiflora
Pseudobombax tomentosum
Annona coriacea
Qualea grandi flora
Diptychandra aurantiaca
Kielmeyera grandiflora
Aspidosperma subincanum
Roupala montana
Eugenia aurata
Casearia sylvestris
Tocoyena formosa
Totals

100

100

Rel.
freq.
0.46
0.46
0.46
0.46
0.46
0.46
0.46
0.46
0.46
0.46
0.46
0.46
100

IV!
1.18
.16
.15
.09
.08
.08
.08
.06
0.99
0.99
0.97 +
0.92
300

TABLE 4. Interfluvial cerrado: species sampled in 100 quadrats (10 x 10m). See Table
for abbreviations.

1.
2
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.

Syagrus comosa
Qualea parviflora
Myrcia albotomentosa
Davilla grandiflora
Qualea grandiflora
Byrsonima verbascifolia
Hymenaea stigonocarpa
Kielmeyera rubriflora
Pouteria ramiflora
Licania humilis
Rourea induta
Vochysia rufa
Andira cuyabensis
Couepia grandiflora
Tabebuia caraiba
Aspidosperma tomentosum
Terminalia brasiliensis
Myrcia uberavensis
Dalbergia miscolobium
Psidium widgrenianum
Mouriri pusa
Salacia crassifolia
Lafoensia pacari
Vatairea macrocarpa
Kielmeyera coriacea
Eugenia punicifolia
Caryocar brasiliense
Mouriri elliptica
Peltogyne confertiflora
Roupala montana
Byrsonima coccolobifolia
Bowdichia virgilioides
Guapira noxia
Annona crassiflora
Andira inermis

Total
ba

Rel.
dens.

Rel.
dom.

355
211
153
152
70
95
50
73
39
36
51
45
33
28
25
32
23
35
19
24
14
27
17
19
18
18
7
18
15
12
14
8
10
11
12

1.952
2.689
1.269
0.736
1.584
0.539
1.210
0.742
1.090
0.881
0.182
0.383
0.532
0.538
0.597
0.295
0.515
0.228
0.483
0.353
0.586
0.209
0.273
0.236
0.158
0.090
0.381
0.075
0.128
0.131
0.087
0.224
0.179
0.143
0.091

18.80
11.18
8.10
8.05
3.71
5.03
2.65
3.87
2.07
1.91
2.70
2.38
.75
.48
.32
.70
.22
.85
.01
1.27
0.74
1.43
0.90
1.01
0.95
0.95
0.37
0.95
0.79
0.64
0.74
0.42
0.53
0.58
().64

9.28
12.78
6.03
3.50
7.53
2.56
5.75
3.53
5.18
4.19
0.86
1.82
2.53
2.56
2.84
1.40
2.45
1.08
2.30
1.68
2.78
0.99
1.30
1.12
0.75
0.43
1.81
0.36
0.61
0.62
0.41
1.06
0.85
0.68
0.43

Rel.
freq.
6.25
5.61
6.56
6.66
4.28
5.04
4.00
4.38
2.95
2.57
3.62
2.47
2.38
2.38
2.00
2.76
1.62
2.09
1.62
1.71
.14
.90
.52
.24
.43
.52
().67
.52
.14
.05
1.14
0.76
0.86
0.95
0.95

IVI
35.02
29.57
20.70
18.21
15.52
12.64
12.39
11.77
10.20
8.66
7.18
6.68
6.66
6.42
6.07
5.86
5.28
5.03
4.92
4.66
4.67
4.33
3.72
3.36
3.13
2.90
2.85
2.83
2.54
2.31
2.30
2.25
2.24
2.21
2.02

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

317

TABLE4(Cont.)
Total
ba

n
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.

Mczilaurus aff. lindaviana

Eriothcca gracilipes
Eugenia aurata
Diospyros coccolobifolia
Eugenia chrysantha
Syagrus petraea
Tabebuia aurea
Pterodon pubescens
Casearia sylvestris
Dimorphandra mollis
Eretnanthus mattogrossensis
Bocageopsis mattogrossensis
Himatanthus phagendaenicus
Ouratea hexasperma
Sclerolobium paniculatum
var. rubiginosum
Miconia albicans
Andira vermifuga
Tabebuia ochracea
Diptychandra glabra
Curatclla americana
Hancornia speciosa
Aspidosperma macrocarpon
Connarus suberosus
Manihot tripartita
Alibertia edulis
Strychnos pscudoquina
Plathymcnia rcticulata
Emmotum nitens
Miconia rubiginosa
Agonandra brasiliensis
Himatanthus obovatus
Xylopia aromatica
Erythroxylum tortuosum
Palicourea rigida
Tocovena formosa
Totals

Rel.
dens.

Rel.
dom.

Rel.
freq.

IVI

-)
10
)
7
(->
c)
5
4
(3
5
5
.?
Li
Ii

0.203
0.114
0.066
0.076
0.089
0.024
0.068
0.098
0.027
0.037
0.021
0.024
0.025
0.012

0.32
0.53
0.48
0.37
0.32
0.48
0.26
0.21
0.32
0.26
0.26
0.16
0.21
0.21

0.96
0.54
0.31
0.36
0.42
0.11
0.32
0.47
0.13
0.18
0.10
0.11
0.12
0.06

0.57
0.76
0.86
0.67
0.57
0.67
0.48
0.38
0.57
0.48
0.48
0.28
0.38
0.38

.85
.83
.65
.40
.31
.26
.06
.06
.02
0.92
0.84
0.75
0.71
0.65

?
i
5
?
?

0.036
0.029
0.015
0.010
0.020
0.057
0.014
0.044
0.011
0.004
0.035
0.033
0.019
0.012
0.005
0.007
0.007
0.006
0.006
0.001
0.001

0.16
0.21
0.16
0.16
0.16
0.05
0.11
0.05
0.11
0.11
0.05
0.05
0.11
0.05
0.05
0.05
0.05
0.05
0.05
0.05
0.05

0.17
0.14
0.07
0.05
0.10
0.27
0.07
0.21
0.05
0.02
0.17
0.16
0.09
0.06
0.05
0.03
0.03
0.03
0.03
0.01
0.01

0.28
0.19
0.28
0.28
0.19
0.10
0.19
0.10
0.19
0.19
0.10
0.10
0.10
0.10
0.10
0.10
0.10
0.10
0.10
0.10
0.10

0.62
0.54
0.52
0.49
0.44
0.42
0.36
0.36
0.35
0.32
0.31
0.31
0.29
0.20
0.20
0.18
0.18
0.18
0.18
0.15
().15

>
>
>
>

1888

21.044

100

100

100

300

TABLE 5. Sandstone cerrado: species sampled by point-centred quarter method in a

systematic grid of 50 points. See Table 1 for abbreviations.

1. Teminalia fagifolia
2 Vochysia petraea
3. Sclerolobium paniculatum
var. subvelutinum
4. Kielmeyera rubriflora
5. Vcllozia flavicans
6. Pterodon pubescens
7. Pouteriarami flora
8. Qualca parviflora
9. Emmotum nitens
10. Davilla grandiflora

Total
ba

Rel.
dens.

Rel.
dom.

Rel.
freq.

IVI

23
41

0.681
0.187

11.50
20.50

23.73
8.17

11.90
17.26

53.14
45.93

21
18
16
8
8
7
7
6

0.341
0.064
0.086
0.181
0.147
0.161
0.100
0.035

10.50
9.00
8.00
4.00
4.00
3.50
3.50
3.00

14.88
2.81
3.74
7.91
6.43
7.01
4.38
1.54

10.71
8.93
6.55
3.57
4.76
2.98
3.57
3.57

36.09
20.74
18.28
15.48
15.19
13.49
11.46
8.11

EDINB. J. BOT. 48(3)

318

TABLE5(Cont.)
Total
ba

Rel.
dens.

Rel.
dom.

Rel.
freq.

IV!

5
5
i\
4
:\
5
I
\
>
>

0.051
0.036
0.010
0.009
0.020
0.015
0.028
0.003
0.023
0.021
0.023
0.014
0.014
0.009
0.007
0.005
0.004
0.004
0.003
0.003
0.003
0.003

2.50
2.50
2.00
2.00
.50
.50
.00
.50
.00
.00
0.50
0.50
0.50
0.50
0.50
0.50
0.50
0.50
0.50
0.50
0.50
0.50

2.23
1.57
0.45
0.39
0.88
0.64
1.23
0.13
1.04
0.93
1.01
0.61
0.61
0.38
0.29
0.22
0.15
0.15
0.13
0.13
0.11
0.11

2.98
2.98
2.38
.79
.79
.79
.19
.79
.19
.19
0.60
0.60
0.60
0.60
0.60
0.60
0.60
0.60
0.60
0.60
0.60
0.60

7.71
7.05
4.83
4.18
4.17
3.92
3.42
3.41
3.23
3.12
2.11
1.71
1.71
1.47
1.39
1.31
1.25
1.25
1.22
1.22
1.21
1.21

200

2.291

n
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.

Micropholis rigida
Byrsonima pachyphylla
Myrcia uberavensis
Connarus suberosus
Heteropterys byrsonimifolia
Syagrus comosa
Byrsonima coccolobifolia
Mimosa xanthocentra
Ilex asperula
Miconia albicans
Caryocar brasilicnse
Norantea goyazensis
Erythroxylum ambiguum
Ferdinandusa elliptica
Ouratca spectabilis
Ladenbcrgia chapadensis
Esenbeckia pumila
Aspidosperma macrocarpon
Eschweilera nana
Bredemeyera altissima
Styrax ferrugineus
Mouriri pusa
Totals

100

100

100

300

6. Synthetic data for the vegetation of the five areas of cerrado studied. n = total
number of individuals; S = number of species; TAD = total absolute density (indiv./ha);
H = average height (m); D = average diameter (cm); TADo = total absolute dominance
(m2/ha); H, = Shannon-Wiener diversity index (with logs, in base 2); J, = Pielou
evenness index (logs, in base 2).
TABLE

Cerrado Area
Murundu
Pebble
Phyllite
Interfluvial
Sandstone

TABLE 7.

nN

TAD

TADo

H,

J,

871
560
240
1888
200

45
65
48
70
32

2713.7
1049.9
1546.4
1888.0
1187.0

2.75
3.59
4.24
3.06
3.38

9.87
11.91
10.39
11.90
10.36

31.96
15.03
16.12
21.04
13.60

3.400
4.899
4.850
4.664
4.057

0.610
0.813
0.868
0.761
0.811

Analytical data for surface horizon (0 20cm) of soils of the five areas of cerrado

studied.
Cerrado Area
Murundu**
Pebble**
Phyllite**
Interfluvial*
Sandstone*

pH in
H,O
4.7
5.5
5.3
4.9
5.2

Chemical Properties
P
K
Ca + Mg
Al
ppm
meq./ lOOg
5.0
10.0
1.2
1.6
0.6

1 10
59
64
13
8

1.0
2.2
1.5
0.3
0.4

1.9
0.5
1.0
0.8
0.2

Parent material: *Sandstone; **Phyllite. O.M.=Organic Material

O.M.

Texture
silt
sand

0/

2.3
2.0
2.0
1.9
0.6

da

0/

47
77
62
87
89

26
7
14
2
2

27
16
24
11
9

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

319

40

20

40

20

X
UJ
Q

40-

20
UJ

OC
UJ

60

40

20

80

6CH

20

UJ

E
Ol
CD

m
u
re
o
re
n

E
o

CD

re
0>

o
re
c

E
o

igno

u
re

o>
re

01

01

u
re

5
nacc

0>

re

TO

re

re
01
o
re

jz
Ol

_Q
O

(/i
>.

re

0)
>

i
<
IT
UJ
I

1-

FIG. 2. Cover value indices of the 16 most important plant families in the five areas of cerrado surveyed.

320

EDINB. J. BOT. 48(3)

important families in CVI, followed by the Sapindaceae (Fig. 2). The typical species are
Magonia pubescens, Callisthene fasciculate Dipteryx alata and Astronium fraxinifolium, which occur commonly as trees of 4-9m, and the shrub Pseudobombax longiflorum
(1.5-2.5m). Other common species are the trees Vatairea macrocarpa, Jacaranda
cuspidifolia, Anadenanthera peregrina var.falcata, Curatella americana and Terrninalia subsericea. The soils present an intermediate texture, higher amounts of Ca + Mg,
K and Al contents and a lower P content. The phyllite bedrock remains usually near or
at the surface (0-40cm in depth).

100-

r1-

8060-

omu

cc

i i

io

osa

20-

mu p e ph in sa
%
100
80
60

11

oin
10

oph

| 1

PD

4020-

in e mu Ph sa

mu

10

ino
pho

100

0
6

omu

60-

ED

4020-

in pe

4 ope

osa

80-

osa

peo

6
2

sa mu

FIG. 3. Polar ordination diagrams (right) and corresponding dissimilarity dendrograms (left) of the five surveyed
areas of cerrado: mu - murundu cerrado; pe - pebble cerrado; ph - phyllite cerrado; in - interfluvial cerrado;
sa - sandstone cerrado. The resulting diagram-dendrogram pairs are based on the following dissimilarity
measures: CC = complemented coefficient of community; PD = percentage difference; ED = euclidian distance.

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

321

The interfluvial cerrado has high values of absolute density and average diameter and
also the highest value of absolute dominance (s Table 6). The Vochysiaceae and
Leguminosae share the highest values of CVI with the families Palmae and Myrtaceae
(Fig. 2). The species with the highest importance value is the dwarf-palm Syagrus
comosa (0.5-1.5m), followed by the low-tree species Qualea parviflora, Qualea grandiflora,Myrcia albotomentosa,Byrsonima verbascifolia and Kielmeyera rubriflora (all
3-5m), and by the shrub species Davilla grandiflora (0.5-3m). Common and taller trees,
of 4-1 lm, are Hymenaea stigonocarpa, Pouteria ramiflora and Licania humilis. The
soils are deep, coarse-textured and with very low levels of P, K and Ca+Mg contents
(Table 7). Although surveyed by different sample sizes, which limits the power of
comparison, the interfluvial cerrado, the phyllite cerrado and the pebble cerrado are
certainly the communities with the highest species diversity (H = 4.6-4.9, see Table 6).
Nevertheless the degree of concentration of individuals in fewer species (given by J,
Table 6) is higher in the interfluvial cerrado.
The sandstone cerrado is less diverse in species than all but the murundu cerrado
(Table 6). It also presents the lowest absolute dominance and a low value of absolute
density. The Leguminosae and the Vochysiaceae share the highest values of CVI with
the Combretaceae (the last based on the presence of just one species: Terminalia
fagifolia) (Fig. 2). The characteristic species are the trees (3-6m), Terminalia fagifolia
and Sclerolobium paniculatum var. subvelutinum and the shrubs (0.5-3.5m) Vochysia
petraea, Kielmeyera rubriflora and Vellozia flavicans. These five species constitute
58.05% of total IVI. As in the interfluvial cerrado, the soils are very sandy and poor, but
present a lower level of organic material and are very shallow: the vegetation occurs
only in the spaces and cracks among the rocks where some soil can accumulate. The
resulting physiognomy is of shrubs and low trees scattered in an irregular pattern
DISCUSSION
The polar ordination diagrams (Fig. 3) show that the murundu cerrado and the sandstone
cerrado represent the extremes of dissimilarity, and that the more the community
structure is emphasized (CC-> PD-> ED diagrams) the more these cerrados are polarized.
In both the polar ordination diagram and the dissimilarity dendrogram based on CC
measures (first pair in Fig. 3; emphasis on floristic composition) the cerrados of the
Baixada Cuiabana (murundu, pebble and phyllite cerrados) form a cohesive group,
separated from the cerrados of the Chapada dos Guimaraes (interfluvial and sandstone
cerrados). In both cases the sandstone cerrado is the most different in terms of floristic
composition. On the other hand, in terms of community structure, which is emphasized
by EDs (lowest pair in Fig. 3), the murundu cerrado is shown quite isolated from the
other cerrados in both the diagram and the dendrogram.
The main floristic dissimilarity between the cerrados of the Chapada dos Guimaraes
(interfluvial and sandstone cerrados) and those of the Baixada Cuiabana (murundu,
pebble and phyllite cerrados) is probably due mostly to differences in soil properties.
The sandstone bedrock of the Chapada dos Guimaraes gives rise to coarse and very

322

EDINB. J. BOT. 48(3)

infertile sediments while the soils of the Baixada Cuiabana are mainly influenced by the
phyllite bedrock, which gives rise to finer and more fertile sediments.
The phyllite cerrado presents many species considered as indicators of the mesotrophic facies cerradao first described by Ratter (1971) in northeastern Mato Grosso and
recognized later in many parts of Central Brazil (Ratter et al, 1973,1977,1978,1988).
This type of cerradao (augmentative of cerrado in Portuguese) occurs mainly on soils
with relatively high levels of Ca and Mg. The authors termed it 'Magonia pubescenslCallisthene fasciculata cerradao', from the two main indicator species which are
also the species with the highest importance values in the phyllite cerrado. Some other
indicator species that could be found in abundance in this cerrado are Pseudobombax
longiflorum, Dipteryx alata, Astroniumfraxinifolium,
Jacaranda cuspidifolia and Anadenanthera peregrina var.falcata. Some species occurring in small numbers - such as
Pseudobombax tomentosum, Acrocomia aculeata, Aspidosperma subincanum, Luehea
speciosa and Guettarda viburnioides - are also absolutely typical of this community.
The pebble cerrado is a widely dispersed type of cerrado that can be found throughout
the Baixada Cuiabana in all sites where the soils are shallow and contain the layer of
pebbles. These aspects favour higher rates of seepage and runoff in the groundwater
regime and also reduce strongly the available volume for root growth and feeding.
Despite having the highest levels of Ca+Mg and P, the apparent richness of these soils
is due mainly to the high proportion of pebbles (c.75% of volume) which are removed
before the analyses. Therefore, the figures in Table 7 really refer to a soil volume about
four times greater than in the other cerrados. The plants therefore have available only
about one-quarter of the apparent Ca+Mg content, which explains the presence of few
indicator species of mesotrophic facies cerradao. Dipteryx alata, Pseudobombax longiflorum and Astroniumfraxinifolium,
however, do occur but only the first is among the
10 species with highest importance values. Four of the most important species are in
the Vochysiaceae: Qualea parviflora, Qualea grandiflora, Salvertia
convallariodora,
and Vochysia rufa. These species, also common in wide areas of cerrado in all Central
Brazil can be considered as amongst the most characteristic species of cerrado in a broad
sense. Curatella americana (Dilleniaceae) which is very common in most cerrados of
the Baixada Cuiabana is usually an indicator of soils with seasonally high water-table
(Oliveira-Filho & Martins, 1986; Oliveira-Filho et al., 1989: Ratter et al., 1988).
The murundu cerrado is a very different community because of the special environmental conditions: the restricted growth-space permanently free of flooding on the
earthmounds, which constitute true ecological islands (Diniz de Araujo Neto et al., 1986;
Oliveira-Filho, 1988). Although the flora of the murundus shows most species in
common with the neighbouring cerrado (the pebble cerrado), its community structure is
very different, with a strong concentration of the importance values in a few species,
particularly Curatella americana. This species, as already mentioned, is an indicator of
seasonal high water levels (floods or seepage) and is also typical of murundus of the
Serra do Roncador (Ratter et al., 1973), of the Ilha do Bananal (Ratter, 1987) and of the
Pantanal Matogrossense (Prance & Schaller, 1982; Ratter et al., 1988). Other cerrado
species which commonly occur in many areas of murundus are Dipteryx alata, Andira

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

323

cuyabensis, Simarouba versicolor and Matayba guianensis. For typical cerrado to occur,
the soils must be freely drained since the majority of species of the cerrado flora do not
tolerate soil waterlogging, even during relatively short periods (Eiten, 1972: OliveiraFilho et al., 1989). So, the cerrado species successfully colonizing the murundus are
those best able to tolerate high soil moisture. The less tolerant species do not succeed
in colonizing the murundus and this seems to happen to the Vochysiaceae whose species
are almost entirely absent in the murundu cerrado despite their abundance in the
neighbouring pebble cerrado. The absence or rarity of the Vochysiaceae were also
noticed by Ratter et al. (1988) in some areas of inundable cerrado of the Pantanal
Matogrossense, although the same author (Ratter et al., 1973) and Oliveira-Filho et al.
(1989) noted Qualea parviflora as characteristic of the cerrado/wet campo transition in
Mato Grosso.
The environmental limiting factors of the sandstone cerrado, at the top of the Chapada
dos Guimaraes represent the other extreme from those of the murundu cerrado. The
vegetation can grow only where some soil accumulates within cracks or in the spaces
among the rocks. The paucity of real soil is aggravated by its coarse texture, by the very
low content of mineral nutrients and organic matter, and by the low water storage
capacity. The species living in such a rigorous environment must have specialized
adaptations. Thus, the species with the highest importance values - Vochysia petraea,
Terminalia fagifolia Sclerolobium paniculatum var. subvelutinum and Vellozia flavicans - are characteristic of drier cerrado locations and are absent or very rare in all the
neighbouring cerrados. However, species such as Kielmeyera rubriflora, Pterodon
pubescens and many others with lower importance values also occur in the interfluvial
cerrado 300m below, at the base of the escarpments.
The interfluvial cerrado also occurs on coarse-textured infertile soils which arose from
the sandstone. Nevertheless, the soils are deeper and with higher water storage capacity
than those of the sandstone cerrado. The environmental limitation of this cerrado is
related principally to water table levels which rise in the rainy season so that water
flushes on the valley sides. The highest level reached by seepage and runoff on the valley
side corresponds to the border line between the cerrado and the marshy campo. The
influence of water-table level within this cerrado is described in details by Oliveira-Filho
etal. (1989).
ACKNOWLEDGEMENTS
We are indebted to the Universidade Federal de Mato Grosso for support received from
many undergraduate students who participated in field activities: in special Silvio A.
Rodrigues, Julio C. Dalponte, Beatriz Schwantes, Cecilia B. Rocha, Susan H. Damasceno, Nelson Laturner, Paulo B. Andrade, and Lyria C. A. Oliveira. We also acknowledge with gratitude the identification of botanical specimens by the taxonomists of the
Universidade Estadual de Campinas and the critical reading of this text by Dr James A.
Ratter, Royal Botanic Garden, Edinburgh, and Dr Paulo S. Oliveira, the Universidade
Estadual de Campinas.

324

EDINB. J. BOT. 48(3)

REFERENCES
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Mato Grosso, Cuiaba.
BROWER, J.E. & Z A R , J.H. (1984). Field and Laboratory Methods for General Ecology.
Dubuque.
CARNEIRO, C.M.R. (1982). A vegetacao dos cerrados: analise e perspectivas. Boletim
Tecnico do Institute Brasileiro de Desenvolvimento

Florestal 7: 5-17.

D I N I Z D E A R A U J O N E T O , M., F U R L E Y , P.A., H A R I D A S A N , M. & J O H N S O N , C.G. (1986).

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Ecology 2: 17-35.
EDIBAP (Empresa de Desenvolvimento Integrados da Bacia do Alto Paraguai. (1979).
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Naturais. 235 pp., Brasilia.
ElDT, R.C. (1968). The climatology of South America, in FTTTKAU, E.J., ILLIES, J.,
K L I N G E , H., S C H W A B E , G.H. & SlOLl, H. Biogeography

and Ecology

in

South

America. The Hague.

EITEN, G. (1972). The cerrado vegetation of Brazil. Botanical Review 38: 201-341.
EVERITT, B. (1980). Cluster Analysis. 136 pp., N. York.
F U R L E Y , P.A., R A T T E R , J.A. & G I F F O R D , D.R. (1988). Observations on the vegetation

of eastern Mato Grosso, Brazil III. The woody vegetation and soils of the Morro de
Fumaca, Torixoreu. Proceedings of the Royal Society of London (B) 235: 259-280
G A U C H , H.G. (1982). Multivariate Analysis in Community Ecology. Cambridge University.
GOODLAND, R. (1971). A physiognomic analysis of the 'cerrado' vegetation of Central
Brazil. Journal of Ecology 59: 411-419.
HOEHNE, F.C. (1923). Phytophysionomia do Estado de Mato Grosso e Ligeiras Notas
a Respeito da Composicdo e Distribuicdo de sua Flora. 104 pp., Secretaria da
Agriculture, S3o Paulo.
MUELLER-DOMBOIS, D. & ELLENBERG, H. (1974). Aims and Methods of Vegetation
Ecology. London, Sydney and Toronto.
OLIVEIRA-FILHO, A.T. (1988). A vegetacao de urn campo de monchoes - microrrelevos
associados a cupins - na regiao de Cuiaba (MT). Doctorate thesis, Univ. of Campinas,
SP.
& M A R T I N S , F.R. (1986). Distribuicao, caracterizacao e composicao floristica das
formacoes vegetais da regiao da Salgadeira, na Chapada dos Guimaraes (MT).
Revista. Brasileira Botdnica 9: 207-223.
, S H E P H E R D , G.J., M A R T I N S , F.R. & S T U B B L E B I N E , W.H. (1989). Environmental

factors affecting physiognomic and floristic variations in an area of cerrado in

Central Brazil. Journal of Tropical Ecology 5:413^31.

POTT, V.J., POTT, A., RATTER, J.A. & VALLS, J.F.M. (1986). Flora da Fazenda
Nhumirim, Nhecolandia, Pantanal. Relacao preliminar. Pesquisa em Andamento
(EMBRAPA CPAP) No. 5, 26 pp.

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

325

G.T. & SCHALLER, G.B.(1982). Preliminary study of some vegetation types of

the Pantanal, Mato Grosso, Brazil. Brittonia 34: 228-251.
RATTER, J.A. (1971). Some notes on two types of cerradao occurring in northeastern
Mato Grosso. /// Simposio sobre o cerrado: 100-102. Universidade de SSo Paulo.
, (1987). Notes on the vegetation of the Parque Nacional do Araguaia (Brazil). Notes
from the Royal Botanic Garden Edinburgh 44: 311-342.
, RICHARDS, P.W., ARGENT, G. & GIFFORD, D.R. (1973). Observations on the
vegetation of northeastern Mato Grosso. I. The woody vegetation types of the
Xavantina-Cachimbo Expedition area. Philosophical Transactions of the Royal
Society of London (B) 226: 449-492.
, ASKEW, G.P., MONTGOMERY, R.F. & GIFFORD, D.R. (1977). Observacoes adicionais
sobre o cerradao de solos mesotroficos no Brasil Central. TV Simposio sobre o
Cerrado: 306-316. Universidade de Sao Paulo.
, , & (1978). Observations on forests of some mesotrophic soils in Central
Brazil. Revista Brasileira Botdnica 1: 47-58.
, POTT, A., VALI, J.P., CUNHA, C.N. & HARIDASAN, M. (1988). Observations on
woody vegetation types in the Pantanal and at Corumba, Brazil. Notes from the Royal
Botanic Garden Edinburgh 45: 503-525.

PRANCE,

APPENDIX
Species recorded in the study areas of the Baixada Cuiabana (BC) and the Chapada dos
Guimaraes (CG)
+, present; -, not observed; br, bromeliad; li, liana; pi, palm; sc, succulent;
sh, shrub; ss, sub shrub; tr, tree
The list contains all species collected and/or observed and not only those recorded in
the sample areas. It is confined to cerrado vegetation (gallery forests and grasslands are
excluded).
growth
form

regions
BC

ANACARDIACEAE
Anacardium humile St. Hil.
Astronium fraxinifolium Schott
A. urundeuva (Fr. All.) Engler
Tapirira guianensis Aublet
T. marchandii Engler
ANNONACEAE
Annona cohacea Mart.
A. crassiflora Mart.
A. dioica St. Hil.
A. pygmaea Warm.
Bocageopsis mattogrossensis R. E. Fr.
Cardiopetalum calophyllum Schldl
Duguetia furfuracea (St. Hil.) Benth. & Hook.
D. lanceolata St. HU.
Xylopia aromatica (Lam.) Mart.
X. sericea St. Hil.

ss
tr
tr
tr
tr
sh
tr
ss
ss
tr
tr
sh-ss
sh-ss
tr
tr

CG

326

EDINB. J. BOT. 48(3)

regions

growth
form
BC
APOCYNACEAE
Aspidosperma cylindrocarpon MiilL Arg.
A. macrocarpon Mart.
A. nobile MiilL Arg.
A. subincanum Mull. Arg.
A. tomentosum Mart.
Hancornia speciosa Gomez
Himatanthus obovatus (M. Arg.) Woodson
H. phagendaenicus (Mart.) Woodson
Odontadenia sp.
Rauwolfia sp.
AQUIFOLIACEAE
Ilex asperula Mart.
I. cerasifolia Reiss.
ARACEAE
Taccarum weddellianum [Brong. ex] Schott
ARALIACEAE
Didymopanax macrocarpum (Cham. & Schldl.) Seem.
D. vinosum (Cham. & Schldl.) Marchand
BIGNONIACEAE
Anemopaegma glaucum Mart.
Arrabidaea brachypoda (DC) Bureau. & K. Schum.
A. corrallina (Jacq.) Sandw.
Cybistax antisyphilitica Mart.
Jacaranda cuspidifolia Mart.
J. decurrens Cham.
Memora pubescens (Sprengel) K. Schum.
Tabebuia aurea Benth. & Hook.
T. caraiba (Mart.) Bureau
T. ochracea (Cham.) Standl.
T. roseoalba (Ridley) Sandw.
Zeyheria digitalis (Veil.) Hoehne
BOMBACACEAE
Eriotheca gracilipes (K. Schum.) A. Robyns
Pseudobombax longiflorum (Mart. & Zucc.) A. Robyns
P. tomentosum (St Hil.) A. Robyns
BORAGINACEAE
Cordia alliodora (Ruiz & Pav.) Oken
C. insignis Cham.
C. glabrata (Mart.) A. DC.
BROMELIACEAE
Ananas sp.
Bromelia balansae Mez
BURSERACEAE
Protium almecega Marchand
P. heptaphyllum (Aublet) Marchand
P. ovatum Engler.
CACTACEAE
Cereus peruvianus (L.) Miller
Melocactus sp.

tr
tr
tr
tr
tr
tr-sh
tr-sh
tr
li
ss
tr
tr
sc
tr-sh
sh
ss
sh
li
tr-sh
tr
ss
ss
tr
tr
tr
tr
sh-ss
tr
tr-sh
tr-sh
tr
ss
tr
br
br
tr
tr
ss
sc
sc

CG

A COMPARATIVE STUDY O F CERRADO AREAS IN BRAZIL

growth
form

327

regions
BC

CG

tr

tr

tr
tr
tr
tr
tr
tr

+
+
-

+
+
+
+
+
+

sh-ss

tr
ss
tr
tr
tr

+
+
+

+
+
+
+
-

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CARDYOCARACEAE
Caryocarbrasiliense Cambess.
CELASTRACEAE
Austroplenckia populnea (Reisseck) Lundell
CHRYSOBALANACEAE
Couepia grandiflora (Mart. & Zucc.) Benth.
Hirtella glandulosa Mart. & Zucc.
H. gracilipes (Hook.) Prance
Iicania hoehnei Pilger
L. kunthiana Hook.
L. humUis Cham. & Schldl.
COCHLOSPERMACEAE
Cochlospennum regium (Mart. & Zucc.) Pilger
COMBRETACEAE
Buchenavia tomentosa Eichler
Combretum discolor Taub.
T. brasiliensis (Cambess.) Eichler
T. fagifolia Mart. & Zucc.
T. subsericeaa Eichler
COMPOSITAE
Eremanthus mattogrossensis Kuntze
Ichthyothere cunabi Mart.
Piptocarpha rotundifolia (Less.) Baker
Vernonia brasiliana (L.) Drude
V. ruficoma Schldl.
CONNARACEAE
Connarus suberosus Planch, var. fulvus (Planchon) Forero
Rourea induta Planch.
DILLENIACEAE
Curatella amencana L.
Davilla elliptica St. Hil.
D. grandiflora St. Hil. & Tul.
EBENACEAE
Diospyros coccolobifolia Mart
ERYTHROXYLACEAE
Erythroxylum ambiguum St. Hil.
E. campestre SL Hil.
E. citrifolium St Hil.
E. cuneifolium (Mart.) E. Schulz
E. deciduum St. Hil.
E. suberosum St. Hil.
E. tortuosum Mart.
EUPHORBIACEAE
Cnidoscolus urens (L.) Arthur
Dalechampia sp.
Manihot tripartita Mull. Arg.
FLACOURTIACEAE
Casearia sylvestris Sw.
GUTTIFERAE
Kielmeyera coriacea (Spr.) Mart
K. grandiflora (Wawra) Saddi

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328

EDINB. J. BOT. 48(3)

regions

growth
form
BC
GUTTIFERAE (contd.)
K. rubriflora Cambess.
HIPPOCRATEACEAE
Peritassa campestris (Cambess.) A.C. Smith
Salacia crassifolia (Mart.) Peyr.
ICACINACEAE
Emmotum nitens (Benth.) Miers
LAURACEAE
Mezilauras crassiramea (Meissn.) Taub.
M. aff. lindaviana Schw. & Mez
Ocotea sp.
LECYTHIDACEAE
Eschweilera nana (Berg) Miers
LEGUMINOSAE CAESALPINIOIDEAE
Bauhinia bongardi Steudel.
B. cuyabensis Steudel.
B. pulchella Mart.
B. rufa Steudel.
Cassia alata L.
C. claussenii Benth.
C. desvauxii (Colladon.) Killip
C. ragosa G. Don.
C. sylvestris Veil.
C. velutina Vogel
Cenostigma gardnerianum Tul.
Copaifera elliptica Mart.
C. langsdorffii Desf.
C. martii Hayne
Dimorphandra mollis Benth.
Diptychandra aurantaca Tul.
D. glabra Benth.
Hymenaea stigonocarpa Mart.
Peltogyne confertiflora (Hayne) Benth.
Sclerolobium aureum (Tul.) Benlh. var. aureum (Tul.) Benth.
S. paniculatum Vogel var. rabiginosum (Tul.) Benth.
S. paniculatum Vogel var. subvelutinum Dwyer
LEGUMINOSAE FABOIDEAE
Acosmium dasycarpum (Vogel) Yakovlev
Andira cuyabensis Benth.
A. inermis Kunth
A. vermifuga Mart.
Bowdichia major (Mart.) Benth.
B. virgilioides Kunth
Centrolobium tomentosum Benth.
Dalbergia miscolobium Benth.
Dioclea tricolor Benth.
Dipteryx alata Vogel
Eriosema rufum E. Mayer
Erythrina velutina Willd.
Galactia glaucescens Kunth
Harpalyce brasiliana Benth.

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329

A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

regions

growth
form
BC
LEGUMINOSAE FABOIDEAE (contd.)
Machaerium acutifolium Vogel
Platypodium elegans Vogel
Pterodon pubescens Benth.
Vatairea macrocarpa Benth.
LEGUMINOSAE MIMOSOIDEAE
Acacia plumosa Lowe
Anadenanthera peregrina (L.) Speg. var. falcata (Benth.)
Altschul
Calliandra parviflora Benth.
Enterolobium contortisiliquum (Veil.) Morong
Inga heterophylla Willd.
Mimosa hirsuta Sprengel
Mimosa pteridifolia Benth.
Mimosa xanthocentra Mart. var. subsericea (Benth.) Barneby
Plathymenia reticulata Benth.
Pithecellobium edwallii Hoehne
Stryphnodendron obovatum Benth.
LOGANIACEAE
Antonia ovata Pohl
Strychnos pseudoquina St. Hil.
LYTHRACEAE
Lafoensia pacari St. Hil.
Physocalymma scaberrimum Pohl
MALPIGHIACEAE
Byrsonima coccolobifolia Kunth
B. coriacea DC.
B. crassa Niedenzu.
B. dealbata Gris.
B. gaultherioides Gris.
B. pachyphylla Juss.
B. subterranea Brade & Mark
B. verbascifolia (L.) Rich, ex Juss.
Heteropterys byrsonimifolia A. Juss.
Heteropterys sp.
MALVACEAE
Pavonia sp.
MARCGRAVIACEAE
Norantea goyazensis Camb.
MELASTOMATACEAE
Miconia albicans (Sw.) Triana
M. fallax DC.
M. ferruginata (DC.) Cogn.
M. guianensis Cogn.
M. rubiginosa (Bonpl.) Tr.
Mouriri clliptica Mart.
M. pusa Gardn
Tibouchina aff. aspera Aublet.
T. pogonanthera (Naud) Cogn.
MELIACEAE
Guarea guidonia (L.) Sleumer

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330

EDINB. J. BOT. 48(3)

growth
form
MENISPERMACEAE
Abuta selloana Eichler
MONJMIACEAE
Siparuna guianensis Aublet
MORACEAE
Brosimum gaudichaudii Trecul
Cecropia cinerca Miq.
C. pachystachya Trecul
Ficus guianensis Aublet
F. cnormis (Mart, ex Miq.) Miq.
MYRISTICACEAE
Virola sebifera Aublet
V. sessilis(A. DC.)Warb.
MYRSINACEAE
Rapanea guianensis Aublet
MYRTACEAE
Campomanesia eugenioides (Cambess.) Legr.
Eugenia aurata Berg
E. bimarginata DC.
E. chrysantha Berg
E. punicifolia (Kunth) DC.
E. uniflora L.
Myrcia albo-tomentosa Cambess.
M. formosiana DC.
M. intermedia (Berg) Kiaersk.
M. aff. nivea Berg
M. pallensDC.
M. suffruticosa Berg
M. uberavensis Berg
M. variabilis DC.
Psidium multiflorum Camb.
P. widgrenianum Berg
NYCTAGINACEAE
Guapira noxia (Netto) Lundell
Neea theifera Oersted
OCHNACEAE
Ouratea floribunda (SL Hil.) Engler
O. hexasperma (St. Hil.) Engler
O. spectabilis (Mart.) Engler
OPEJACEAE
Agonandra brasiliensis Miers
PALMAE
Allagoptera campestris (Mart.) Kuntze
A. leucocalyx (Drude) Kuntze
Acrocomia aculeata (Jacq.) Lodd. ex Mart.
Astrocaryum campestre Mart.
Butia leiospatha (Mart.) Becc.
Syagrus comosa (Mart.) Becc.
S. petraea (Mart.) Becc.
POLYGALACEAE
Bredemeyera altissima Benth.

regions
BC

CG

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A COMPARATIVE STUDY OF CERRADO AREAS IN BRAZIL

growth
form

regions
BC

POLYGONACEAE
Coccoloba mollis Casar.
C. cf. cordifolia Meissner
PROTEACEAE
Roupala montana Aublet
RHAMNACEAE
Rhamnidium elaeocarpum Reisseck.
RUBIACEAE
Alibertia concolor (Cham.) K. Schum.
A. edulis Rich.
A. sessilis (Cham.) K. Schum.
A. verrucosa Moore
Chomelia ribesioides Benth.
Coussarea hydrangeaefolia Benth. & Hook.
Ferdinandusa elliptica Pohl
Guettarda viburnioides Cham. & Schldl.
Ladenbergia chapadensis Moore
Palicourea coriacea (Cham.) K. Schum.
P. rigida Kunth
Rudgea viburnioides (Cham.) Benth.
Sabicea cana Hook.
Tocoyena formosa (Cham. & Schl.) K. Schum.
RUTACEAE
Escnbeckia pumila Pohl
Zanthoxylum arenarium Engl.
Z. cinereum Engl.
SAPINDACEAE
Dilodendron bipinnatum Radlk.
Magonia pubescens St. Hil.
Matayba guianensis Aublet
Serjania erecta Radlk.
S. sp.
Talisia subalbens Radlk.
SAPOTACEAE
Chrysophyllum marginatum (Hook. & Am.) Radlk.
Micropholis rigida Pierre
Pouteria lateriflora (Benth.) Radlk.
P. ramiflora (Mart.) Radlk.
SIMAROUBACEAE
Simaba trichilioides St. Hil.
Simarouba versicolor St. Hil.
SMILACACEAE
Smilax sp.
SOLANACEAE
Solanum lycocarpum St. Hil.
STERCULIACEAE
Guazuma tomentosa Kunth
G. ulmifolia Lam.
Helicteres sacarolha St. Hil.
Waltheria sp.

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331

CG

332

EDINB.J.BOT.48(3)
growth
form

regions

BC
STYRACACEAE
Styrax camporum Pohl
S. fertugineus Nees & Mart.
TTLIACEAE
Luehea speciosa Willd.
ULMACEAE
Trema micrantha Blume
VELLO23ACEAE
Vellozia flavicans Mart.
VERBENACEAE
Aegiphila sellowiana Cham.
Vitex cymosa Bert.
VTTACEAE
Cissus sp.
VOCHYSIACEAE
Callisthene fasciculata (Spr.) Mart
Qualea grandiflora Mart
Q. multiflora Mart.
Q. parviflora Mart.
Salvertia convallariodora St Hil.
Vochysia divergens Mart.
V. haenkeana Mart.
V. petraea Warm.
V. pyramidalis (Spr.) Mart.
V. rufa (Spr.) Mart.
V. tucanorum (Spr.) Mart.

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CG